A replacement name for Sphenomorphus keiensis (Kopstein, 1926) from the southeastern Moluccas, Indonesia (Reptilia: Squamata: Scincidae) with a redescription of the species

G.M. Shea & J.P. Michels

Shea, G.M. & Michels, J.P. A replacement name for Sphenomorphus keiensis (Kopstein, 1926) from the southeastern Moluccas, Indonesia (Reptilia: Squamata: Scincidae) with a redescription of the species. Zool. Med. Leiden 82 (52) 31.xii.2008: 737-747, fi gs 1-2.— ISSN 0024-0672. Glenn M. Shea, Faculty of Veterinary Science, B01, University of Sydney, NSW 2006, Australia, and Research Associate, The Australian Museum, 6 College St, Sydney, NSW 2000, Australia (gshea@mail. usyd.edu.au). J. Pieter Michels, Research Associate, National Museum of Natural History, P.O. Box 9517, 2300 RA Leiden, The Netherlands.

Key words: Scincidae; Sphenomorphus; systematics; nomenclature; Indonesia. Lygosoma keiensis Kopstein, 1926 is demonstrated to be a junior primary homonym of Lygosoma cyano- gaster keiensis Sternfeld, 1918. The junior homonym, now placed in Sphenomorphus, is given the replace- ment name S. capitolythos, and redescribed from the holotype, the only known specimen.

Introduction

Many homonyms exist among nominal species-group taxa described in the scin- cid genus Lygosoma, the genus to which most skinks were referred during the second half of the 19th and fi rst half of the 20th century (Duméril & Duméril, 1851; Boulenger, 1887; De Rooij , 1915). This is particularly the case among species from southeast Asia and the Australopapuan region. Many of these homonyms are secondary, arising from Boulenger’s (1887) combination in Lygosoma of species originally described in a number of genera, and a variety of replacement species names were created at the time (Boulenger, 1887; Ahl, 1925). However, some are primary homonyms, refl ecting the long period aft er Boulenger’s monograph when most new skink species were re- ferred to Lygosoma. Lygosoma was subsequently split into numerous genera, most notably by Smith (1937), Mitt leman (1952) and Greer (1974). For example, the 112 species referred to Lygosoma in the most recent monographic treatment of the skinks of the Indonesian/ Papuan archipelago (De Rooij , 1915) are currently assigned to 16-17 genera (there is recent disagreement about the validity of the genus Apterygodon; Mausfeld & Schmitz, 2003). The dismemberment of Lygosoma, and the lack of any complete checklist of names in the family since Boulenger (1887), has resulted in litt le att ention being paid in recent years to the existence of primary homonyms among species originally described in Lygosoma but now in diff erent genera. Most of these homonyms apply to names now considered synonymous with other taxa and not in present use. Therefore there is no need to provide them with a substitute name (Article 23.3.5; International Commission on Zoological Nomenclature, 1999). However, there still remain some problematic names. 738 Shea & Michels. A replacement name for Sphenomorphus keiensis. Zool. Med. Leiden 82 (2008)

One publication particularly rife with homonyms is Sternfeld (1918), oft en incor- rectly cited as published in 1920 (e.g., Loveridge, 1948; Tyler, 1968; Zweifel, 1979; Mys, 1988; Brown, 1991). Sternfeld described 16 new species and subspecies of skinks in this paper. Eight of these are homonyms. Sternfeld twice proposed the same species or sub- species name for two taxa in the same paper: Lygosoma (Hinulia) maindroni wolfi (p. 395) and Lygosoma (Otosaurus) wolfi (p. 397), and Lygosoma (Riopa) albofasciolatum boett geri (p. 418) and Lygosoma (Emoa) boett geri (p. 406), possibly considering that either the diff ering subgeneric assignment, or the allocation of names to subspecies rather than species, avoided the homonymy. These primary homonyms were identifi ed by Mertens (1924), who proposed the nomina nova Otosaurus sternfeldi (currently Sphenomorphus concin- natus (Boulenger, 1887)) for L. wolfi , and Riopa albofasciolatum poehli (currently Eugongy- lus albofasciolatus (Günther, 1872)) for L. a. boett geri. Three of Sternfeld’s skinks, Lygosoma (Keneuxia) smaragdinum nigrum (p. 400), Lygo- soma (Emoa) cyanogaster aruensis (p. 405) and Lygosoma (Hinulia) jobiense elegans (p. 397), were junior secondary homonyms at the time of description, respectively of Eumeces niger Jacquinot & Guichenot, 1853) (now Emoia nigra, see Brown, 1991), Eumeces aruensis Doria, 1874 (now Sphenomorphus jobiensis (Meyer, 1874), see Boulenger, 1887 and Zweifel, 1980), and Hinulia elegans Gray, 1838 (the current identity of which is unknown (Cogger et al., 1983), although from the illustration subsequently provided by Gray (1845), it may be a synonym of Eulamprus tenuis (Gray, 1831)). Lygosoma jobiense elegans is also a junior primary homonym of Lygosoma elegans Boulenger, 1897 (now Lobulia elegantoides (Ahl, 1925) and a junior secondary homonym of Euprepes (Tiliqua) elegans Fischer 1883 (now Lygosoma fernandi (Burton, 1836)) and itself a junior primary homonym of Euprepes elegans Peters, 1854, long considered to be part of the genus Mabuya Fitzinger, 1826, but more recently implicitly tranferred to the resurrected genus Euprepis Wagler (see Maus- feld et al., 2002), and still more recently to Trachylepis Fitzinger, 1843 (see Bauer, 2003) with recognition that Euprepis was not available for the Afro-Malagasy lineage formerly in Mabuya; Fischer (1884) proposed the replacement name Euprepes leoninus for Euprepes (Tiliqua) elegans. Mertens (1929) provided the substitute name Dasia smaragdinum melas (now in Lam- prolepis) for the secondary homonym L. smaragdinum nigrum Sternfeld, 1918, although the recognition of subspecies within Lamprolepis smaragdinum has mostly been avoided for over half a century (Kinghorn, 1928; Burt and Burt, 1932; Hediger, 1933, 1934; Greer, 1970; McCoy, 1980; Mys, 1988; Crombie & Pregill, 1999), and neither Sternfeld’s nigrum nor Mertens’ substitute name has subsequently appeared in the literature. Should the Nissan Atoll Lamprolepis be accorded taxonomic recognition in the future, Sternfeld’s nigrum is to be used as this name is no longer homonymous. The homonymy of Sternfeld’s elegans was noted by Loveridge (1948: 346), although as no author since Sternfeld has applied the name to a species or subspecies distinct from Sphenomorphus jobiensis, no substitute name is needed. Sternfeld’s aruensis is currently placed in the synonymy of Emoia longicauda (Ma- cleay, 1877) (see Brown, 1991), and again no substitute name is needed. An eighth skink taxon described by Sternfeld (1918) is also a homonym, though this time as a senior primary homonym, and this homonymy has not been previously rec- ognised, nor is any subsequently published name available for the junior homonym. Sternfeld (1918) described Lygosoma (Emoa) cyanogaster keiensis, while Kopstein (1926) Shea & Michels. A replacement name for Sphenomorphus keiensis. Zool. Med. Leiden 82 (2008) 739 described Lygosoma (Homolepida) keiensis, both from the Kei Islands (now Kai Islands, or Kepulauan Kai, Indonesia). The epithet is formed incorrectly in both instances, as the name Lygosoma is neutral in gender. The epithet keiensis is to be corrected to keiense ac- cording to Articles 31.2 and 34.2 of the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature, 1999). This mandatory action has no further implications for the primary homonymy observed. Sternfeld’s subspe- cies is currently placed in the genus Emoia, while Kopstein’s species is currently placed in Sphenomorphus. The purpose of this paper is to provide a replacement name for Kopstein’s junior homonym, and a more thorough description and diagnosis of the species, still known only from the holotype.

Systematics

Lygosoma (Emoa) cyanogaster keiense Sternfeld, 1918

L[ygosoma]. [(Emoa)] cyanogaster keiensis Sternfeld, 1918: 405; Mertens, 1922: 175. Lygosoma [(Emoa)] cyanogaster, Kopstein, 1926: 90. Emoia cyanogaster, Loveridge, 1948: 366. [Emoia] keiensis, Mitt leman, 1952: 25. Emoia cyanogaster keiensis, Brown, 1954: 264; Mertens, 1967: 74. Emoia longicauda, Brown, 1991: 49; How et al., 1998: 134.

Holotype.— SMF 15334 (Mertens, 1967: 74). Type locality.— “Kei-Inseln, ? Molukken […] Langgur (Kei)” (Sternfeld, 1918: 405). Current status.— Emoia longicauda (Macleay, 1877). Discussion of citation history.— Kopstein (1926) was the fi rst to comment on Stern- feld’s subspecies, stating that all of the Moluccan specimens he had collected agreed with Sternfeld’s diagnosis of L. c. aruense. No material from the Kai or Aru Islands was available to him, but nevertheless he remarked that the purported diff erences in col- oration and patt ern were untenable in view of the great chromatic variation within Lygosoma species. Consequently, he referred his specimens to a monotypic L. cya- nogaster. Loveridge (1948) rejected both aruense and keiense as well, but erroneously stated that Sternfeld had not provided any diagnosis. Mitt leman (1952) listed the spe- cies-group taxon keiense in combination with Emoia, but this action did not refl ect an intent to elevate it to species status for he noted “[I]n no sense does this list purport to be a checklist of the forms considered valid.” Brown (1954) left the taxonomic status of the subspecies indeterminate, though restricting E. c. cyanogaster (Lesson, 1826) to the Solomon Islands, and recognising E. c. longicauda (Macleay, 1877) for populations in New Guinea. Mertens (1967) gave as current status Emoia cyanogaster keiensis without reference to a source, although presumably following Brown’s listing. Brown (1991) in his monographic revision of the genus, placed Sternfeld’s subspecies (misspelt twice as E. c. keinensis) in the synonymy of E. longicauda (Macleay, 1877), raising the latt er to species status, although noting that further material was needed to resolve the status of the Aru and Kei Island populations. How et al. (1998), reporting on material from the Aru Islands, followed Brown’s (1991) synonymy of Sternfeld’s subspecies by not using trinominals in E. longicauda. 740 Shea & Michels. A replacement name for Sphenomorphus keiensis. Zool. Med. Leiden 82 (2008)

Sphenomorphus capitolythos, nomen novum for Lygosoma keiense Kopstein, 1926 (fi gs 1-2)

Lygosoma [(Homolepida)] keiensis Kopstein, 1926: 86. L[ygosoma]. (Sph[enomorphus].) keiensis, Brongersma, 1942b: 156. [Sphenomorphus] keiensis, Greer & Parker, 1967: 19. Sphenomorphus keiense, Scott et al., 1977: 12; Whitaker et al., 1982: 47; Welch et al., 1990; Monk et al., 1997: 435.

Holotype.— RMNH 5088, an ethanol-preserved specimen of undetermined sex, col- lected March 1923, by Dr Felix Kopstein. Type locality.— “Elat, Gross-Kei” (Kopstein 1926: 86). In contemporary geographi- cal nomenclature, this locality is Elat (or Banda Elat) on Kai Besar, Kepulauan Kai, Maluku Tenggara Province, Indonesia (5°39’S 132°39’E). Diagnosis.— A medium-sized species of Sphenomorphus (SVL 81 mm) with short limbs, widely separated when adpressed. The combination of grooved subdigital la- mellae, a scaly lower eyelid lacking a central window, four supraoculars, third pair of chin shields medially separated by three scales but in lateral contact with the infrala- bials, no postsupraocular scale, and a temporal region with no fragmentation or divi- sion of the last two supralabial scales, single primary temporal scale or upper and lower secondary temporal scales, and with the upper secondary temporal overlap- ping the lower secondary temporal, will diff erentiate this species from all other mem- bers of the Sphenomorphus group of lygosomine skinks (Greer, 1979a) in Indonesia, the New Guinea region, and Australia.

Fig 1. Dorsal view of the holotype of Sphenomorphus capitolythos, nom. nov.

Fig 2. Dorsal, right lateral and ventral views of the head of the holotype of Sphenomorphus capitolythos, ▶ nom. nov. with (inset) detail of abnormal temporal region on left side of head (dashed line represents normal extent of primary temporal scale). Scale bar = 2 mm. Shea & Michels. A replacement name for Sphenomorphus keiensis. Zool. Med. Leiden 82 (2008) 741 742 Shea & Michels. A replacement name for Sphenomorphus keiensis. Zool. Med. Leiden 82 (2008)

Description of holotype.—Nomenclature of head scales follows Taylor (1935), with exceptions as noted. Head scalation.— Nasals moderately separated, nostril slightly posteroventrally lo- cated in nasal; supranasals and postnasals absent; prefrontals large, broadly separated medially; frontoparietals in broad contact; interparietal with parietal eye at junction of middle and posterior third; parietals in contact behind interparietal; nuchals 2/3; a single scale between upper secondary temporal and anterior nuchal on each side; supraoculars four, fi rst two in contact with frontal; supraciliaries nine; posteriormost supraciliary (= anterior pretemporal of Greer, 1983) extends medially behind fourth supraocular, to contact frontoparietal on right side, but not on left side; posteriormost supraciliary not in contact with postocular, which instead wedges dorsally between penultimate supra- ciliary and uppermost postsubocular (= posterior pretemporal of Greer, 1983); single anterior and single posterior loreal, each with height and length similar, but posterior loreal dorsally a litt le longer than tall; presuboculars four; suboculars one; postsubocu- lars fi ve; lower eyelid scaly, lacking a transparent window; supralabials seven, fi ft h be- low centre of eye, but separated from it by subocular series, sixth and seventh undivided; postsupralabials two; primary temporal single; secondary temporals two, lower over- lapped by upper (on left side, the primary temporal is abnormally small, and the lack of its ventral part results in the last supralabial extending anteriorly to contact the postocu- lar series; the arrangement of the scales of this region on the right side is typical of other sphenomorphine skinks with undivided temporal scales); external auditory meatus round, maximum diameter a litt le smaller than height of eye, tympanum deeply re- cessed; anterior margin of external auditory meatus lacking lobules; infralabials seven, fi rst two in contact with postmental; three pairs of transversely enlarged chin chields, all laterally contacting the infralabial series; fi rst pair of chin shields in medial contact; sec- ond pair separated by a median scale; third pair separated by three median scales. Body and limb scalation.— Body scales in 31 rows at midbody (32 slightly anterior to this level); paravertebral scales not broader than adjacent dorsal scales, 59 from ante- riormost nuchal to last scale anterior to level of hindlimbs (63 to level of posterior mar- gin of hindlimbs); all scales polished and glossy; at high magnifi cation, dorsal and lat- eral scales with many fi ne low longitudinal striations, but ventral scales smooth; me- dian pair of preanals enlarged, overlapping more lateral preanals; lamellae below fourth toe 16, with a postaxial groove distinguishing a larger anterior and smaller pos- terior portion; scales above fourth toe six basally, three at middle of toe, reducing to two distally, with only terminal scale single; a sharp demarcation between rounded granu- lar scales on sole and imbricate dorsal scales. Measurements.— Snout-vent length 80.5 mm; axilla-groin interval 44 mm; tail dis- tally regenerated; forelimb length 15 mm; hindlimb length 22 mm; head length 14.9 mm; head width 11.3 mm; head depth 8.8 mm. Osteology (based on a radiograph of the type, and on direct examination of the dentition). — Presacral vertebrae 26, fi rst three lacking ribs; sacral vertebrae two; tail regenerated from 17th postsacral vertebrae; ribs 6-8 sternal, 9-11 mesosternal, last four ribs short; phalangeal formula of manus and pes 2.3.4.5.3 and 2.3.4.5.4 respectively; premaxillary teeth nine (sum of left and right sides); maxillary teeth 18/17; postorbital bone possibly absent (the radiograph does not show a clearly distinct element in this region, although the postfrontal, jugal and squamosal are distinguishable). Shea & Michels. A replacement name for Sphenomorphus keiensis. Zool. Med. Leiden 82 (2008) 743

Coloration.— Colour of the recently preserved specimen (aft er at most three years in preservative) was described by Kopstein (1926) as dorsally brown with irregularly arranged dark fl ecks, half a scale in size. Sides of the neck with faded grey reticulations. Lateral surfaces lighter brown. Ventral side uniform yellowish. The coloration in 2006, 80 years later, was similar, but paler. The dark fl ecks are located on both dorsal and lateral surfaces of body and tail, and are mostly located in the centre of scales, but oc- casionally obliquely oriented to cross scale rows. The upper and lower labial scales are dark-edged.

Discussion

Kopstein remarked that his species was most closely related to L. unilineatum De Rooij , 1915. The latt er taxon was recently reassigned to Eugongylus Fitzinger, 1843 by Greer and Shea (2000). However, S. capitolythos is not assignable to Eugongylus, based on supradigital scalation (multiple rows of scales in S. capitolythos vs. a single row in Eugongylus), nuchal scale orientation (obliquely to parietals vs. fl ush with parietals), preanal scales (median pair enlarged and overlapping adjacent lateral preanal scales vs all preanals more or less equal in size, with median pair overlapped by adjacent lateral preanal scales), number of presacral vertebrae (26 vs 28-30) and supranasals (absent vs usually present). In the fi rst three of these character states, and in possessing nine pre- maxillary teeth, the species is clearly a member of the Sphenomorphus group of Greer (1979a). However, within this major lineage, the generic assignment of this species is less clear. In most scalational features, and in coloration, the species is similar to Glap- hyromorphus nigricaudis (Macleay, 1877), from which it diff ers in having deeply grooved subdigital lamellae (vs smoothly rounded, ungrooved) and a greater number of para- vertebral scales (63 vs 52-58, see Greer, 1979b). In the former feature, it is more similar to Sphenomorphus species occurring in New Guinea (Smith, 1937; Shea & Greer, 1999). However, it diff ers from the most speciose group in this region (the S. maindroni group, Greer & Shea, 2004) in lacking a postsupraocular scale, although in similarity with the postsupraocular of the S. maindroni group, the terminal supraciliary (or anterior pre- temporal) extends to the frontoparietal (but on one side only). There has been no rigorous morphological or biochemical analysis of relationships within the Sphenomorphus group of lygosomine skinks that has included both Glaphyro- morphus and members of the S. maindroni group. The genus Sphenomorphus itself is cur- rently undiagnosed in terms of derived character states within the Sphenomorphus group, and is likely to consist of more than one lineage on the basis of morphological variation. Similarly Glaphyromorphus is currently undiagnosed in terms of derived char- acter states within the Sphenomorphus group, and represents a morphotype rather than a lineage (Greer, 1989, 1990). DNA sequence evidence for the polyphyly of Glaphyromor- phus has been presented by Reeder (2003). Reeder also reported a major monophyletic lineage consisting of all Australian genera within the Sphenomorphus group, although his sampling of non-Australian representatives of the group was limited to just three species (representing Prasinohaema, Scincella and the Sphenomorphus muelleri species group). Hence, it is possible that his Australian lineage may include some non-Austral- ian taxa, including some species currently assigned to Sphenomorphus. Given the present uncertainty about the limits and relationships of Sphenomorphus and Glaphyromorphus, we tentatively retain S. capitolythos in Sphenomorphus. 744 Shea & Michels. A replacement name for Sphenomorphus keiensis. Zool. Med. Leiden 82 (2008)

Etymology

The specifi c epithet is a noun in apposition and honours Felix Kopstein (born 4 June 1893 in Vienna, Austria-Hungary [Adler stated Austria, but at that time Austria was part of the Habsburg Austro-Hungarian dual monarchy]; died 14 April 1939 in The Hague, the Netherlands), a physician and author of the last review of the Moluccan reptilian fauna (Adler, 1989). Capitolythos is derived from caput (Latin for ‘head’) and lythos (Greek for ‘stone’), in reference to the two probable German compound words in the name Kop- stein, German for head and stone respectively. This unconventional form of patronym was chosen because of the existence of Lygosoma emigrans kopsteini Brongersma (1942a). The species emigrans is presently assigned to the genus Glaphyromorphus (Greer, 1990) but the subspecies has not been re-evaluated in subsequent studies of the genus in the Lesser Sundas (Auff enberg, 1980; Greer, 1990; Aplin et al., 1993), and has only been men- tioned in a listing of Brongersma’s named taxa by Hoogmoed (1995).

Acknowledgements

Dr Marinus Hoogmoed (RMNH) kindly loaned the holotype of the species to the senior author for examination. Thanks are also extended to Mr Bertus van Tuij l, Dr Isaäc J.H. Isebrücker, Dr Axel Groenveld and Ms Elsbeth Zwart (ZMA) for granting access to the library and providing working space to the junior author. Dr Mieke Konings (Am- sterdam) most kindly assisted in assessing the correctness of the name capitolythos.

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Received: 21.viii.2008 Accepted: 19.ix.2008 Edited: L.P. van Ofwegen